In
population genetics, 'genetic load' or 'genetic burden' is a measure of the cost of lost alleles due to
selection ('selectional load') or
mutation ('mutational load'). It is a value in the range
, where 0 represents no load.
The concept was first formulated in 1937 by
JBS Haldane, independently formulated, named and applied to humans in 1950 by
H. J. Muller[1], and elaborated further by Haldane in 1957.
[2]
Definition
Genetic load is the reduction in selective value for a population compared to what the population would have if all individuals had the most favored genotype.
[3] It is normally stated in terms of
fitness as the reduction in the mean fitness for a population compared to the maximum fitness.
Mathematics
Consider a single
gene locus with the
alleles
, which have the
fitnesses
and the
allele frequencies respectively. Ignoring
frequency-dependent selection, then genetic load (
) may be calculated as:
:
where
is the
maximum value of the fitnesses
and
is
mean fitness which is calculated as the mean of all the fitnesses weighted by their corresponding allele frequency:
:
where the
allele is
and has the fitness and frequency
and
respectively.
When the
, then (1) simplifies to
:
Causes of genetic load
Load may be caused by selection and mutation.
Mutational load
Load caused by mutations is known as mutational load.
Selectional load
Selection occurs when the fitnesses of particular alleles are
inequal, hence selection always exerts a load.
With
directional selection, the allele frequencies will tend towards an equilibrium position with the fittest allele reaching a frequency in
mutation-selection balance. As mutations are rare, this is effectively fixation. Consider two alleles
and
. If
, then at equilibrium,
and
, hence
, and
.
If the mean fitness is 0, the load is equal to 1, but the population goes
extinct.
Segregational load
In contrast to directional selection,
heterozygote advantage always exerts a load at equilibrium.
Creationist criticism
Some
creationists (such as
Henry M. Morris) have suggested that mutational load would increase over time and thus make populations inviable. However, they ignore the effect of selectional load acting to weed out (decrease frequency of) deleterious mutations.
References
1.
2. The cost of natural selection, JBS Haldane, , , Journal of Genetics, 1957
3. Some possibilities for measuring selection intensities in man, JF Crow, , , Hum. Biol, 1958
External links
★
''The Cost of Natural Selection Revisited'', Leonard Nunney, Ann. Zool. Fennici 40:185-194 (pdf file)
★
Genetic load, from ''Evolution A-Z'' by Mark Ridley
★
"Understanding Genetic Load" - a paper by Dr. L. Monroe
★
Creationist Claim CB120: Genetic load from ''An Index to Creationist Claims'' by Mark Isaak